Life-stages, exploitation status and habitat use of Lutjanus goreensis (Perciformes: Lutjanidae) in coastal marine environments of Lagos, SW Nigeria
Ciclo de vida, estado de explotación y hábitat de Lutjanus goreensis (Perciformes: Lutjanidae) en ambientes marinos costeros de Lagos, suroeste de Nigeria
Kafayat, A. Fakoya1*, Martins, A. Anetekhai1, Shehu, L. Akintola1, Abdulwakil, O. Saba1 & Mikhail A. Abass2*
Abstract
The Gorean snapper, Lutjanus goreensis is an important component of artisanal fisheries and trawl landings in the Gulf of Guinea. Despite its economic importance, there is a dearth of information on size structure and life history strategies of the species. Therefore, the objectives of this study were to provide baseline data on the life stages, exploitation status and habitat use for the species in Nigeria. Monthly samples were obtained from artisanal and trawl catches in Five Cowrie Creek and Lagos coastal waters between December 2008 and December 2010, respectively. Length-frequency distributions of the fishes caught were analysed to provide preliminary information on mean and modal lengths at capture and life - history strategies based on habitat use and estuarine-dependency for L. goreensis. A total of 822 specimens of L. goreensis were collected from Five Cowrie Creek while 377 specimens were collected from Lagos coastal waters. Total length varied between 7.90-34.90cm for creek samples and from 21.90-56.10cm for marine samples. Length-frequency histograms showed polymodal size distributions in creek and marine samples. Length-frequency distributions of L. goreensis showed a high abundance of juveniles (<20cm) and sub-adults (20-35cm) which accounted for 84.1% and 68.4% of creek and marine samples examined, respectively. For the creek samples, fish in modal length class of 13.00-13.99cm were the most exploited while in the marine samples, length classes of 29.00-30.99cm and 31.00-32.99cm constituted the most frequently exploited fishes. Increase in total lengths from the creek (mean±SD; 16.19±3.73cm) to the marine habitat samples (32.89±6.14cm) indicated ontogenetic shift in habitat use. Occurrence of a predominant juvenile population in Five Cowrie Creek by L. goreensis suggests estuarine-dependency and is indicative of a temporary juvenile habitat or a migratory corridor. In conclusion, data from the presently reported study and previous studies demonstrated that juvenile L. goreensis displays estuarine dependency and habitat flexibility. Hence, this underscores the importance of preserving estuarine environments as essential fish habitats to prevent overfishing. The study also concludes that the species is vulnerable to recruitment overfishing in the marine environment especially as a consequence of shrimping. Consequently, it advocates for ban on all fishing activities during peak spawning periods in breeding grounds and shrimp ground assemblage.
Resumen ]]>
El pargo Lutjanus goreensis es un componente importante de la pesca artesanal y de arrastre en el Golfo de Guinea. A pesar de su importancia económica, hay una escasez de información sobre la estructura de tamaño y las estrategias de historia de ]]>
L. goreensis. Un total de 822 ejemplares de L. goreensis se obtuvieron de Five Cowrie Creek, mientras que 377 muestras de las aguas costeras de Lagos. La longitud total varió entre 7.90-34.90cm para muestras del estuario y 21.90-56.10cm para marinas. ]]>
L. goreensis mostraron una alta abundancia de juveniles (<20cm) y subadultos (20-35cm) que representaron el 84.1% y el 68.4% de los estuarios y las muestras marinas examinadas respectivamente. Para las muestras del estuario, peces de talla 13.00-13.99cm fueron los más explotados, mientras ]]>
L. goreensis sugiere una ]]>
L. goreensis presentan una dependencia a los estuarios y una flexibilidad en el uso de hábitat. Por lo tanto, esto subraya la importancia de preservar los ambientes estuarinos como hábitats esenciales de peces para evitar la sobrepesca. El estudio también concluye ]]>
Palabras clave:Lutjanus ]]>
, distribución de frecuencia del largo, uso de hábitat, estapas de vida, dependencia estuarina, status de explotación.
Lutjanidae popularly known as snappers comprises a large family of the order Perciformes, important ]]>
Lutjanus goreensis (Valenciennes, 1830) is a medium to large and deep-bodied species of snappers (over 35cm) (Martinez-Andrade, 2003) common to 50cm but capable of attaining maximum total length of 80cm (Allen, 1985). It belongs to the ]]>
Population structure ]]>
Spatial separation of size-classes of fish species suggests movement from one habitat to another with ontogeny (Cocheret de la Morinie`re, Pollux, Nagelkerken, & van der Velde, 2003). Ontogenetic migrations have been inferred from spatial separation of size - classes in many fish species including lutjanids ]]>
Lutjanus argentimaculatus (Sheaves, 1995); L. jocu (Moura, Francini-Filho, Chaves, Minte-Vera, & Lindeman, 2011); L. argentiventris (Aburto-Oropeva, Dominguez-Guerrero, Cota-Nieto, & Plomozo-Lugo, 2009); L. apodus and L. griseus (Hammerschlag-Peyer, ]]>
The concept of “estuarine dependence” acknowledges that fish use of estuarine and oceanic habitats is a continuum. An estuarine-dependent species is defined as one in which estuaries, or similar habitats, are the principal environments for at least part of the life cycle and without which a viable population would cease to exist (Nagelkerken, & van der ]]>
Notwithstanding its economic importance, studies related to size structure, life-stages, habitat ]]>
L. goreensis in the estuarine and marine environments are either poorly documented or lacking. Globally, several species of shallow and deepwater snapper stocks are vulnerable to over-fishing with serious consequences for conservation and management (Grandcourt, Abdessalaam, & Franklin, 2006). Though, exploitation status of the snapper species is suggested to be over-exploited in Nigerian coastal waters ]]>
L. goreensis. Available data from previous studies of Fakoya, Abass, Owodeinde, Lawson, and Ojo (2010); Orhibhabor and Ogbeibu (2010) in estuarine habitats consist of low sample sizes.
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Therefore, this study is unique in that it provides the first data on the life-stages, exploitation status and habitat use of the Gorean snapper, L. goreensis from comparative analysis of the length-frequency distributions in estuarine and marine environments, respectively. Thus, the objectives of this study are to give information on the mean and modal lengths at capture, to describe ]]>
L. goreensis.
Materials and Methods
Study sites: This study was carried out in Five Cowrie Creek and Lagos coastal waters, South-West, Nigeria as shown in Fig. 1. Five Cowrie Creek is one of the numerous adjoining creeks to the Lagos Lagoon. It is connected to the Lagoon at two ends; the Lagos Habour which opens to the inshore (coastal) waters off Lagos ]]>
Casuarina equisetifolia, Terminalia catappa and sparse occurrence of grasses Chromolaena odorata (Adekanmbi, & Ogundipe, 2009).
The fishing grounds of Lagos ]]>
2 which is located between the high-water mark on the shore and lower limit of the thermocline (Ssentongo, Ukpe, & Ajayi, ]]>
Fish sampling: Fish samples were purchased from licensed trawlers (cod-ends with stretch mesh size of 44mm and 76mm of shrimp and fish trawl nets, respectively operating in the marine coastal waters of Lagos. Artisanal fishermen using hook and line exclusively were the sources of fish samples from Five Cowrie Creek and also additional samples from marine coastal waters, respectively. Fish specimens were identified with the aid of taxonomic ]]>
Size-distributions, life-stages, ]]>
Samples of L. goreensis from the creek and Lagos coastal waters were measured (total length, TL to the nearest 0.1cm) using the traditional fish measuring board and weighed (whole body weight to the nearest 0.1g) with an electronic balance (Mettler PM 400). Each fish specimen was cut open and sex determined by observing the gonads as there is no sexual dimorphism in this species. To minimize systematic error due to collection from commercial catches, ]]>
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The life stages (adult, sub-adult and juvenile) of L. goreensis were determined based on total length. Juvenile and sub-adult life stages were based on size classes of L. argentiventris (Aburto-Oropeva et al., 2009); L. apodus and
L. griseus (Hammerschlag-Peyer et al., 2013) as: small-sized or juveniles (<20cm); medium-sized or sub-adults (20-30cm). The large-sized or adult life-stage was estimated based on size at first sexual maturity which was defined as 43% of maximum length for shallow-water and continental snapper species (Grimes, 1987). The maximum length for the species was based on a maximum total length of 80cm in the Gulf of Guinea for the species (Allen, 1985). Hence, all fish less than 35cm but greater than 30cm were also included as sub-adults in this study. This size-class of >35cm corresponded to large-adults in
L. peru (Reddy et al., 2013).
Mean and modal lengths at capture were estimated for the species in the two habitats to assess exploitation status of the species. Length-frequency distributions of sampled fishes in the two habitats were used to infer habitat use. The species was considered to undergo possible ontogenetic habitat shifts based on mean total length. If the mean total length of a fish species ]]>
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Data analyses were performed by IBM SPSS statistics version 20. The data on total lengths were analysed using descriptive statistics and expressed as mean ± standard deviation, mode and percentage. Student’s t-test (α=0.05) was used to assess for significant differences in mean total lengths: (i) between creek and marine samples of Lutjanus goreensis
and (ii) between trawl caught fish and artisanal caught fish.
Results
Size-distributions and life-stages: Information relating to size distributions and life- stages of creek ]]>
L. goreensis are given in Table 1. A total of 822 specimens were sampled from the creek while 377 specimens were examined from the coastal waters. Total lengths of creek samples ranged from 7.90cm to 34.70cm and body weight ranged from 9.51g to 695.60g whereas composite ranges for marine samples varied between 21.90cm and 56.10cm total length and from 156g to 2 975g body weight. Juveniles ]]>
In marine samples, fish length ]]>
Lengths at capture: The length-frequency histograms showed polymodal distributions and percentage frequency of different size classes in samples of
L. goreensis from the creek and coastal waters. Creek samples caught with hook by the artisanal fishery were largely unsexed and had a mean of 16.19cm ±3.73 standard deviation. By comparison, marine samples caught with the same gear had a mean of (35.54±8.08cm) (Table 1), indicating significant variations in sizes of the fish species in the two ecological habitats.
In creek samples modal class of 13.00-13.99cm with a frequency of 12.4% was the most exploited (Fig. 2A). Length classes of ≤9.00cm and 34.00-34.99cm with frequencies of 0.2% and 0.1% respectively, accounted for the least exploited fishes. Composite marine samples had a mean length of 32.89±6.14cm ]]>
Table 1). Modal class of 29.00-30.99cm was the most exploited with a frequency of 15.1% followed closely by the length class of 31.00-32.99cm with a frequency of 14.9% (Fig. 2B). Approximately, 58.1% of total sample was <33.00cm thus representing a majority of size classes fished below the modal class. Males (n=189) ranged from 21.90 to 56.10cm with a mean of 32.30±6.59cm while females (n=188) ranged from 22.60 to 47.60cm with a mean of 33.47±5.60cm (
Table 1). There was no significant difference in mean total lengths observed between males and females (Student t-test, p>0.05). In males and females, specimens of 29.00-30.99cm and 31.00-32.99cm were also the most exploited (Fig. 2B).
Analysis by gear revealed that the ]]>
Table 1). The Student’s t-test indicated a significant difference in the mean total lengths of trawl caught fish and artisanal caught fish (p<0.05) with larger sizes occurring in the latter. Modal classes of 29.00-30.99cm and 31.00-32.99cm represented the most exploited fishes with equivalent frequencies of 15.0% by trawl fishery. In artisanal fishery, the modal class of ]]>
Fig. 2C).
Habitat use: Minimum sizes in the marine habitat (21.90cm) were always larger than minimum sizes in the estuarine habitat (7.90cm), indicating that the smallest individuals of the species occur only in the creek and not in the marine habitat. ]]>
Mean size of the fish was significantly higher in the marine habitat (32.89±6.14cm) than in the creek (16.19±3.73cm) (Student’s t-test, p<0.5) showing clear ontogenetic shifts in habitat use by L. goreensis. The species also showed estuarine-dependence in its juvenile stage. Juvenile life-stage of the species (<20cm) was the most dominant (84.1% of total sample) in the creek and showed gradual movement over size range of 20-35cm to the marine habitat where the species occurred only as sub-adults and adults.
Discussion
The largest size of the species from Five Cowrie Creek was much larger than the largest fish previously reported in other studies for Nigeria (Lawson, 2005; Orhihabor, & Ogbeibu, 2010; 2012), Ghana (Aheto et al., 2011; Okyere, Aheto, & Aggrey-Fynn, 2011); Bamboung, Senegal (Faye, Le Loc’h, Thiaw, & Tito de Moraïs, 2012) and Gabon (Mamonekene et al., 2006). ]]>
L. goreensis noted in the present study and previous studies is as a result of the larger sampling size collected in this study, differential gear selectivity – hook and line in this study against gillnet (stretched mesh size of 10-50mm) in Gabon (Mamonekene et al., 2006); pole seine net (stretched mesh size 5mm) in Ghana (Okyere et al., 2011); fishing pressure and lengths of sampling period. These factors, notably sampling size and gear selection, ]]>
The present study indicated that juveniles consisted of the most dominant life-stage in the creek while ]]>
L. goreensis from estuarine/brackishwater environments as juveniles (Lawson 2005; Orhihabor, & Ogbeibu, 2010, 2012), in coastal wetlands systems of Ghana (Aheto et al., 2011; Okyere et al., 2011) and in Complex of Protected Areas of Gabon (Mamonekene et al., 2006). Results of the ]]>
Analysis of mean, ]]>
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Appearance of adult sizes in trawl and artisanal catches suggested that the species probably moves to shallower waters near the coast where the fisheries occur. García-Contreras, Quiñónez-Velázquez, Morán-Angulo, & Valdez-Pineda (2009) observed a similar shoreward migration by the Amarillo snapper, L. argentiventris. A plausible reason for the onshore migration could be that in Nigerian ]]>
L. analis (Manjarrés-Martínez, Gutiérrez-Estrada, Mazenet-González, & Soriguer, 2010). Heavy rains in Nigeria occur between May and September, often causing the sea to be rough. Another factor could be that the peak spawning period for lutjanids coincides with maximum rainfall and ]]>
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This study also indicated that there was no restriction on the sizes of juveniles and sub-adults fished from the creek. Mean and modal total lengths, respectively suggests that the fish is prone to growth overfishing such that replenishment by adult stock in the marine habitat declines. It is important to restrict fishing of the species in the creek because they are prominently juveniles.
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In the present study, the size and age at first maturity of Lutjanus goreensis or related species in the Eastern central Atlantic are not known. Therefore, the mean total length and the most fished length class of marine samples could be below the maturation size for the species, the consequence of which would be recruitment overfishing. Sub-adults sampled from the marine ]]>
Comparison between length-frequency distributions of L. goreensis also suggests that an increase in mean total lengths from the creek to the marine habitat denotes habitat ]]>
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Two migration patterns are suggested in this study. First, a mono- or uni-directional migration pattern from the creek to the marine habitat in which fish size increases. The strategy employed by the species appears to be that of an ontogenetic shifter where habitat use is dependent on life-stage as described by Jaxion-Harm, Saunders and Speight (2012). Gonad development and dietary shifts are known theoretically to initiate ontogenetic migrations in fishes (Gillanders, Able, Brown, Eggleston, & Sheridan, 2003; Moura et al., 2011; Jaxion-Harm et al., 2012). In ]]>
L. goreensis from the creek to the marine habitat where may be an abundance of highly diverse spectrum of preferred preys exist. Similarly, Cocheret de la Moriniere et al. ]]>
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Predominance of sub-adults in marine samples lends anecdotal evidence that sub-adult staging areas for the species were located within the larger marine habitat. This is consistent with the views of Gillanders et al. (2003). From the foregoing, ecological conditions required for sexual maturation of L. goreensis were obviously absent in ]]>
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Secondly, the probability of a cross- or bi-directional migration between the creek and the marine habitat may also be inferred from the concurrent size classes of sub-adults in both habitats. Feeding may be a fundamental reason for the marine sub-adult population of the species to return to the creek where there may be preferred small preys. Similarly, Okyere et al. (2011); Aheto et al. (2011) suggested occasional migration of adult marine fish species (interpreted here to include sub-adults forms) to estuaries where they once lived as juveniles for the purpose of ]]>
L. griseus and L. analis (Claro et al., 2009). Adult-sized L. griseus and mullids also migrate daily into seagrass/mangrove habitats which serve as juvenile habitats for feeding (Honda et al., 2013). Thus, it may be assumed that based on the observations of sub-adults in both habitats, the species is a habitat generalist. ]]>
Occurrence of a predominant juvenile population in the creek is indicative of a juvenile habitat. The postulated life cycle in medium to large species in subfamily Lutjaninae (Martinez- Andrade, 2003) and movement of snapper larvae in Barrier Lagoon complex and Strand Coasts (Ezenwa et al., 1990), both insinuate migration of early juveniles to the estuarine environment from the coastal waters where they occurred as larvae, while spawning most apparently must have occurred in the marine habitat. Data from the ]]>
L. goreensis in marine samples strongly suggests that coastal waters are clearly not a nursery ground for the species.
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In addition, the paucity of juveniles in the marine habitat clearly demonstrates that juvenile phase of L. goreensis in the creek has a clear dependency on the estuarine environment for its survival. The species spends only part of its life cycle in the creek. Martinez-Andrade (2003) also considered L. goreensis as an estuarine-dependent species for its juvenile phase; Aheto et al. (2011); Okyere et al. (2011); Brochier, Ecoutin, de Morais, Kaplan, and Lae (2013) considered L. goreensis as a marine species accessory in estuaries implying that it resides in estuaries ]]>
The ability to utilise multiple estuaries as putative juvenile habitats and later to reassemble as adults in larger groups in the marine habitat is suggestive of the evolution of meta-population structures, a trait of marine fish species which exhibit estuarine dependency (Vasconcelos, 2009). Both tidal creeks and mangroves are habitats of juvenile snappers (Harbone et al., ]]>
L. goreensis occurring in the mangrove-depauperate tidal creek such as Five Cowrie Creek, Omogoriola et al. (2012) had reported their occurrence in mangroves of the Eastern parts of Lagos Lagoon.
A mangrove-poor habitat has a ]]>
Otherwise, by virtue of being one of the two routes channelling water from the Lagos Lagoon Complex into ]]>
In conclusion, the species is more of an ontogenetic shifter than a habitat generalist. Estuarine dependency and habitat flexibility are characteristics of the juvenile ]]>
L. goreensis underscores the importance of preserving estuarine environments as essential fish habitats for juveniles of commercially important fish species. Subtle evidence suggests that large reproductive adults may be more vulnerable to exploitation during the rainy season while biodiversity loss ]]>
L. goreensis in the marine environment, there is need to identify spawning period of the species and shrimp ground assemblage for proper ]]>
Acknowledgments
We thank the Chief technologist, ]]>
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1. Department of Fisheries, Faculty of Science, Lagos State University, Ojo, Lagos State Nigeria; kafade@yahoo.com, kafayat.fakoya@lasu.edu.ng, anetekhaimartins@gmail.com, drsilfa@yahoo.com, shehu.akintola@lasu.edu.ng ,sabaola@gmail.com
2. Nigerian Institute for Oceanography and Marine Research, 3 Wilmot Point, Off Bar Beach Road, Victoria Island, Lagos State, Nigeria; mikailabass@yahoo.com
Received 06-V-2014. Corrected 25-IX-2014. Accepted 24-X-2014.